N400 (neuroscience)

The N400 is a component of time-locked EEG signals known as event-related potentials (ERP). It is a negative-going deflection that peaks around 400 milliseconds post-stimulus onset, although it can extend from 250-500 ms, and is typically maximal over centro-parietal electrode sites. The N400 is part of the normal brain response to words and other meaningful (or potentially meaningful) stimuli, including visual and auditory words, sign language signs, pictures, faces, environmental sounds, and smells.

The N400 was first discovered by Marta Kutas and Steven Hillyard in 1980. They conducted the first experiment looking at the response to unexpected words in read sentences, expecting to elicit a P300 component. The P300 had previously been shown to be elicited by unexpected stimuli. Kutas and Hillyard therefore used sentences with anomalous endings (i.e.I take coffee with cream and dog), expecting to see a P300 to the unexpected sentence-final words. However, instead of eliciting a large positivity, these anomalous endings elicited a large negativity, relative to the sentences with expected endings (i.e. He returned the book to the library) In the same paper, they confirmed that the negativity was not just caused by any unexpected event at the end of a sentence, since a semantically expected but physically unexpected word (i.e. She put on her high-heeled SHOES) elicited a P300 instead of negativity in the N400 window. This finding showed that the N400 is related to semantic processing, and is not just a response to unexpected words.

The N400 is characterized by a distinct pattern of electrical activity that can be observed at the scalp. As its name indicates, this waveform peaks around 400 ms post-stimulus onset, with negativity that can be observed in the time window ranging from 250-500 ms. This latency (delay between stimulus and response) is very stable across tasks—little else besides age affects the timing of its peak. The N400 is a negative component, relative to reference electrodes placed on the mastoid processes (the bony ridge behind the ear), and relative to a 100 ms pre-stimulus baseline. Its amplitude can range from -5 to 5 microvolts. However, it is important to note that in studies using the N400 as a dependent measure, the relative amplitude of the waveform compared to another experimental condition (the "N400 effect") is more important than its absolute amplitude. The N400 itself is not always negative—it is just a more negative-going deflection than that seen to other conditions. Its distribution is maximal over centro-parietal electrode sites, and is slightly larger over the left side of the head for visual words, although the distribution can change slightly depending on the eliciting stimulus.

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